Panurgine bees are diverse and abundant in temperate areas of the Americas but poorly represented to nearly absent in the tropics. We describe and illustrate five distinctive new species of the genus Protandrena that occur at high altitudes (20003400 m) in the Andes, from Venezuela to Ecuador. The species are also described to make the names available in forthcoming papers on their biology. These Andean species resemble some members of the subgenus Heterosarus but differ from it, as well as from any other subgenera of Protandrena, primarily in characters of the male genitalia and hidden sterna. The South American Protandrena s. l. are morphologically highly diverse and a complete study of the group is needed before supraspecific names are proposed for unusual species. Thus, to avoid further nomenclatural changes, we decided not to place these species in a new subgenus or any of the available subgenera. We also provide notes on the biology for some of the species.

The purposes of this work are to describe and illustrate five distinctive new species of the bee genus Protandrena Cockerell from the higher tropical Andes (20003400 m), and to make their names available in forthcomings papers on their biology. Panurgine bees are diverse and abundant in temperate areas of the Americas but poorly represented to nearly absent in the tropics (Michener 2000). As far as we know, only a few species in the genera Acamptopoeum Cockerell and Protandrena occur in the northwestern part of South America, from Venezuela to Ecuador (Gonzalez 2004; Gonzalez & Engel 2004). The Protandrena species described in this work, as well as at least three other presumably related species from the Peruvian and Bolivian Andes, have the following characteristics in common: body predominantly dark brown to black, two submarginal cells (Fig. 1), propodeal triangle basally glabrous, and mesoscutum finely punctured. The sixth sternum of the male has a V-shaped midapical emargination (Figs. 1316), and the tibial scopa of the female is sparsely covered with simple or minutely branched hairs. Thus, in the key to the subgenera of Protandrena of Michener (2000), these species run out to the subgenus Heterosarus Robertson. However, they differ from Heterosarus and any other known subgenera of Protandrena primarily in male characters of the hidden sterna and genitalia. The apical lobes of the seventh sternum are narrow and distally retrorse (Figs. 1720) as in Protandrena s. str. and Metapsaenythia Timberlake, but the apex of these lobes lack the distinctive spatulate hairs present in the latter. In addition, Metapsaenythia has a hairy propodeal triangle and a metasoma frequently red. The gonostylus and gonocoxite are hairless and completely fused (Figs. 2532). Such structures have long branched hairs and can be distinguished, at least ventrally, in the subgenera Heterosarus and Metapsaenythia.

Given these peculiarities, our first impression was to place these Andean species into a new subgenus. However, the South American Protandrena s. l. are morphologically highly diverse, and many species cannot easily assigned to any of the known subgenera (e.g., Ruz 1986; Ruz & Chiappa 2004; Ramos & Melo 2006). A complete revision of the group is needed before supraspecific names are proposed for unusual species. Therefore, to avoid further nomenclatural changes, we decided not to place these Andean species in a new subgenus or in any of the available subgenera. We also provide notes on the biology for some of the species.

MATERIAL AND METHODS

The morphological description and illustrations were made using an Olympus SZ60 stereomicroscope. Morphological terminology follows that of Michener (2000) and, in general, Harris (1979) for surface sculpturing. Description format follows that of Gonzalez (2004). The abbreviations F, S, T, OD, and PD are used for antennal flagellomere, metasomal sternum and tergum, and ocellar diameter and puncture diameter, respectively. Type specimens are deposited in the following institutions: Instituto de Ciencias Naturales (ICN), Universidad Nacional de Colombia (Bogotá), and Entomology Division, Natural History Museum, University of Kansas (SEMC), Lawrence, KS, USA.

Protandrena bachuesp. nov.

Diagnosis. Males can be recognized by the following combination of characters: clypeus with an inverted T-shaped yellow maculation (Figs. 1, 2), antennal flagellum strongly crenulate (Fig. 5), and S5 with midapical margin bearing a long row of spines (Fig. 9). Females can be distinguished by the combination of facial fovea about 2.4x longer than broad (Fig. 33) and antennal flagellum weakly crenulate, especially the basal segments (Fig. 35).

Etymology. The specific name refers to the Muisca goddess Bachué. The Muisca was a South American nation that lived mainly in modern-day Cundinamarca and Boyacá highlands of Colombia. According to the legend, Bachué came out from the lake of Iguaque with a boy in her arms, giving origin to the human race. The Lake of Iguaque is located in the Iguaque Sanctuary, the type locality of this species.

Floral records. The pollen from the scopae of a single female specimen collected on December 2001 in Chingaza Natural Park (Cundinamarca, 4º 30 N, 73º 40'N, 3150 m) was removed and kindly examined by G. Bogotá. Most of the pollen grains (~ 90 %) belonged to Gaultheria sp. (Ericaceae), a common genus of shrubs with showy red or white flowers. The remaining pollen types were, in order of abundance, as follows: Vallea stipularis L.F. (Elaeocarpaceae), Loutergia-Plagiocheilus (Asteraceae, tubiflorae group), and Oxalis sp (Oxalidaceae). Given the relatively low abundance of these pollen grains in the sample, they may be contaminants or were gathered accidentally during nectar visits.

Protandrena maximina sp. nov.

Diagnosis. Females of this species are similar to P. rangeli sp. nov. in the unmodified antennal flagellum (as in Fig. 36) and the shape of S6 (Fig. 42). However, P. maximina sp. nov. has a longer facial fovea (0.7x vs. half of scape length) and is much larger than P. rangeli sp. nov. (head width, 1.92.1 vs. 1.61.7 mm).

Etymology. This species honors Prof. Maximina Monasterio at the Universidad de los Andes (Mérida, Venezuela), for all her efforts in promoting the study and conservation of high Andean ecosystems.

Protandrena wayrurongasp. nov.

Diagnosis. This species is similar to P. bachue sp. nov. but differs by the shape of the clypeal yellow maculation (Fig. 3), shape of mandible (Fig. 8), number and distribution of midapical spines on S5 (Fig. 10), and pubescence on S6 (Fig. 14).

Comments. We also examined a single female collected on XI-16 1970 from Puracé, a locality in Colombia [Departamento of Cauca, 2º15'N, 76º25'W; 3400 m] about 200 km north of Papallacta. The facial foveae and slightly modified antennae of this specimen are similar to the female of P. bachue sp. nov. but it is smaller, about the size of P. wayruronga sp. nov. This specimen, deposited in SEMC, might be the female of the latter species but we did not designate it as a paratype.

Etymology. The specific epithet means "bee" in the Quechua language. Quechua is the main spoken language by several indigenous populations of the Ecuadorian highlands, where this bee occurs.

Protandrena rangeli sp. nov.

Diagnosis. The male of this species can be distinguished by its small body size, unmodified antennal flagellum (Fig. 6), clypeus with a yellow maculation as in Fig. 4, and S5 with a row of four spines on the midapical margin (Fig. 11). The female of P. rangeli sp. nov. resembles that of P. maximina sp. nov. but it is smaller and has longer facial fovea than the latter (see also comparative comments on that species).

Holotype: , COLOMBIA, Boyacá: Santuario de Fauna y Flora de Iguaque, Quebrada Carrizal, 2830 m, I-8 1992. C. Sarmiento (ICN). Paratypes: 3, 7, one male and two females with the same data as the holotype; four females from the same locality but on the following dates and collectors: I-5 1992, VIII-21 1998, D. Forero; VII 2000, P. Reina; and two males from Cundinamarca, Bogotá, Universidad Nacional de Colombia, 2600 m, VIII-15 1999, V. Gonzalez (ICN, SEMC).

Etymology. This species honors Prof. Orlando Rangel (ICN) for his efforts in promoting the study and conservation of the floral diversity in Colombia. In addition, Prof. Rangel pioneered palynological research in the Andes.

Biological notes. One of us (VG) collected males of this species on Veronica persica Poir (Scrophulariaceae) and Oxalis sp. (Oxalidaceae) from gardens in the Universidad Nacional de Colombia, Bogotá. Also, on June 23 2000, VG found unemerged males and females inside their cells along a ditch next to the visitor center in Iguaque. The cells were approximately 4.1 mm in diameter and were located individually at the end of short (about 17 mm) vertical tunnels that were filled with coarse soil. Individual nest entrances could not be found.

Protandrena guarnensis sp. nov.

Diagnosis. The species can be recognized by the combination of the following characters: antennal flagellum unmodified, long, in repose reaching the mesoscutellum in the male (Fig. 1); body dark reddish brown, without yellow marks in both sexes; face distinctly depressed, especially in the male, above alveolus toward ocellocular space. This depression resembles the antennal depression of Chilicola (Hylaeosoma) (Colletidae, Xeromelissinae). Other species closely related to P. guarnensis sp. nov. from Ecuador, Peru, and Bolivia have different combination of characters and their genitalia structure and associated sterna are quite different.

Etymology. This specific epithet refers to the type locality of this species, Guarne, a beautiful rural area in the northwestern Andes of Colombia.

Acknowledgments. This paper is dedicated to Robert Brooks, who urged VG to describe and study these bees since 1999. We want to thank C. D. Michener for continuous advice and support. We also thank D. Bennet for providing the photographs and J. Pinzón for illustrations of P. bachue sp. nov. and P. rangeli sp. nov.; G. Bogotá and J. L. Fernández (ICN) for kindly identifying the pollen and plant samples; M. S. Engel and Z. Falin for access to bee specimens at SEMC, and L. Packer and C. Rasmussen for their comments and suggestions that helped improve this note. The University of Kansas (KU), Undergraduate Program in Biology, Department of Ecology and Evolutionary Biology, KU General Research Fund and US-Israel Binational Science Foundation grant 2000-259 (to D. Smith & Y. Lubin) provided financial support for VG through teaching assistantships and laboratory facilities.